Cerebellum and Eyeblink Conditioning
The cerebellum and brainstem constitute the essential neural circuit responsible for the acquisition and expression of the classically conditioned eyeblink response in numerous mammalian species, including humans. In this simple form of motor learning, a
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Derick H. Lindquist, Joseph E. Steinmetz, and Richard F. Thompson
Abstract
The cerebellum and brainstem constitute the essential neural circuit responsible for the acquisition and expression of the classically conditioned eyeblink response in numerous mammalian species, including humans. In this simple form of motor learning, a neutral conditioned stimulus (CS) overlaps and coterminates with a mildly aversive unconditioned stimulus (US), resulting, eventually, in the production of an eyeblink conditioned response (CR) to the CS alone. The forebrain is engaged when this basic delay procedure is made more difficult – for instance, if the CS and US are separated by a brief stimulusfree gap of time. In either case, it is generally accepted that the critical memory trace is formed and stored in the cerebellar interpositus nucleus (IP). The cerebellar cortex also plays a key role in normal acquisition by modulating the amplitude and/or timing characteristics of the eyeblink CR. Owing to the welldefined nature of the neural circuit, and the close correspondence between animal and human studies, eyeblink conditioning has been successfully used to investigate cerebellar dysfunction across a variety of human populations. Herein, research related to three representative disorders is discussed: fetal
D.H. Lindquist (*) Department of Psychology, The Ohio State University, 1835 Neil Ave., Room 49, 43210 Columbus, OH, USA e-mail: [email protected] J.E. Steinmetz Department of Psychology, The Ohio State University, 230 N. Oval Mall, 43210 Columbus, OH, USA e-mail: [email protected] R.F. Thompson Department of Psychology, University of Southern California, 3641 Watt Way HNB-522, 90089 Los Angeles, CA, USA e-mail: [email protected] M. Manto, D.L. Gruol, J.D. Schmahmann, N. Koibuchi, F. Rossi (eds.), 1175 Handbook of the Cerebellum and Cerebellar Disorders, DOI 10.1007/978-94-007-1333-8_50, # Springer Science+Business Media Dordrecht 2013
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D.H. Lindquist et al.
alcohol syndrome, attention-deficit hyperactivity disorder, and schizophrenia. The results advance understanding of these and similar clinical pathologies and the cerebellar deficits that may underlie them.
Introduction Over the last half century, a number of simple model systems have been developed that allow the brain correlates of learning and memory to be systematically explored, permitting learning-related behavioral changes to be linked to specific neural circuitry and experience-dependent alterations in synaptic strength. Pavlovian or classical conditioning, which utilizes simple contingency-paired learning with easily measured forms of acquired or conditioned responding, has proven particularly amenable to neurobiological analyses. For instance, in delay eyeblink conditioning (EBC), a neutral conditioned stimulus (CS) is paired and overlaps with an eyeblink-eliciting unconditioned stimulus (US), such as a puff of air directed at the cornea or a mild periorbital electrical shock. The US produces a vigorous reflexive eyeblink termed the unconditioned
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